Japanese honeysuckle (Lonicera japonica) An Alien Plant Report Prepared by: Forest Starr, Kim Martz and Lloyd Loope United States Geological Survey Biological Resources Division In cooperation with: American Water Works Association Research Foundation Maui County Board of Water Supply March, 1999 Japanese honeysuckle (Lonicera japonica) What is Japanese honeysuckle? Why is it bad? Japanese honeysuckle is a fast growing vine with glossy green leaves, and fragrant white to yellow flowers. Japanese honeysuckle’s twining stems strangle vegetation and anything else in its way. In Koke’e on Kaua’i, Japanese honeysuckle is spreading and considered by some as having the potential to be worse than Banana poka. On Maui, Japanese honeysuckle is being used as a landscape plant. Please do not plant Japanese honeysuckle. If you see this plant in the wild, pull it up and alert the HEAR project. What should you do if you see this plant? 1. Call the Hawaii Ecosystems at Risk (HEAR) project at 572-4418. 2. Please do not plant this in your yard. 3. Pull out the vine, if possible. Otherwise, cut near the root of the plant, and treat with an herbicide. 4. For more information or additional copies of this flyer, call 572-4418, or websurf to www.hear.org. You can help keep Maui no ka oi. United States Geological Survey Biological Resources Division Japanese honeysuckle ( Lonicera japonica) Maui County Board of Water Supply 2 American Water Works Association Research Foundation Japanese honeysuckle (Lonicera japonica) Overview: Japanese honeysuckle is an invasive plant in Hawaii, New Zealand, and the eastern part of North America. It has been widely used in horticulture and is known to escape from gardens into adjacent lands, climbing and smothering other plants in its path. It has been outlawed in both parts of the United States and New Zealand. In Hawaii, even with its invasive history, it is currently sold and planted as ground cover. Plants in Hawaii were originally thought not to produce fruit, yet viable seed producing plants have been found on Maui in the course of this study. Japanese honeysuckle has been observed spreading vegetatively from gardens into adjacent lands. Its ability to produce viable seeds allows for long distance dispersal by birds or other animals which would allow the plant to invade many more areas currently free of Japanese honeysuckle. Japanese honeysuckle may be just beginning to occupy its potential range. Potential public involvement: The invasive characteristics and problems associated with Japanese honeysuckle have been well documented in the eastern United States and New Zealand where it has been banned through noxious weed acts. Though, despite the literature which warns of Japanese honeysuckle’s invasiveness, it is currently being planted widely in Hawaii. This information may not be reaching the general public or it does not seem like a real threat. Perhaps, only those acutely aware of alien species problems realize the threat Japanese honeysuckle poses to Hawaii’s natural communities and watersheds, even with the large amounts of information that has been written documenting its invasiveness. Getting this information out to the general public, to nursery owners, to landscapers, to children in schools, on the television, and on the radio to raise public awareness about the potential harm of Japanese honeysuckle could decrease its popularity and decrease the use of the plant in landscaping. Ultimately, making it illegal to possess, as New Zealand and many parts of the United States have done, would be the best way to minimize further spread. Common name: Japanese honeysuckle, honekakala (Wagner et al. 1990), Chinese honeysuckle (Williams and Timmins 1997). Latin name: Lonicera japonica Thunb. Taxonomy: In the Caprifoliaceae (honeysuckle) family, Lonicera is a genus of more than 150 species of shrubs and woody twining climbers. Found in North America and Eurasia south to northern Africa, the Philippines, and southwestern Malesia (Wagner et al. 1990). Description: This vigorous climber grows up to 30 ft (9 m), has glossy, dark green leaves. Pairs of fragrant white flowers, ageing yellow or sometimes purple tinged (Turner 1997). Japanese honeysuckle ( Lonicera japonica) 3 Sprawling and twining lianas; young stems pubescent. Leaves ovate, elliptic, oblong, or broadly lanceolate, blades 3-8 cm long, 1-3.5 cm wide, pubescent, becoming glabrate above, entire or young lower leaves sometimes lobed. Flowers 2 in axillary cymes, bracts 1-2 cm long, bracteoles suborbicular, ca. 1 mm long; corolla white, turning yellowish or tinged pink, 2-lipped, 2-3 cm long. Berries bluish black, globose, 6-7 mm in diameter (Wagner et al. 1990). Noxious weed acts: Japanese honeysuckle is not listed on the United States Federal Noxious Weed or Seed Lists. Japanese honeysuckle is not on the Hawaii Noxious Weed List. Japanese honeysuckle is categorized as an exotic weed under the Illinois Exotic Weed Control Act of 1987. As such, commercial sale in Illinois is prohibited (Nyboer 1990). In New Zealand, in 1995, Japanese honeysuckle was included on the Forest Friendly list of plants unsuited for planting because of its known weediness (Craw 1994). It is no longer permitted to be sold, propagated, distributed or commercially displayed on the basis that it is now a Regional Surveillance Plant Pest within the Auckland Region (Auckland Regional Council 1997). Native range: Native to Asia (Wagner et al. 1990). Range of invasion: Becoming a serious weed in areas such as eastern North America, New Zealand, and Hawaii. Other areas where Japanese honeysuckle has invaded and is now widespread include the southern parts of Australia through New South Wales and Victoria, where it is regarded as a very serious threat to native vegetation (Carr et al. 1992), and wastelands in parts of Southern Chile (Williams and Timmins 1997). In Hawaii, the first naturalized collection of Japanese honeysuckle was made in 1951 (Wagner et al. 1990). Today, Japanese honeysuckle is widespread in Hawaii (Cronk and Fuller 1995). In Hawai’i, it is escaping cultivation in cool, mesic to wet areas in Koke’e State Park, Kaua’i, on O’ahu, East Maui, and scattered localities on Hawai’i (Wagner et al. 1990). On Maui, Japanese honeysuckle has been widely planted, though few naturalized or wild populations exist far from original plantings. At most of these wild populations, Japanese honeysuckle is moving vegetatively from intentional plantings into adjacent lands. One wild population is located in a Eucalyptus grove with no signs that it had been planted. It is not known whether this population has established through secondary spread, by bird or other animal, from a nearby planting or whether is an old house site and the Japanese honeysuckle is being persistent. Japanese honeysuckle may just be beginning to establish in the wild on Maui. In New Zealand, Japanese honeysuckle was being sold for horticulture use in 1872 (Esler 1988), and was first collected from the wild in 1926. Habitats invaded in New Zealand include shrublands, forest margins, disturbed or secondary forest, wetland margins, coastal areas and inshore islands, roadsides, farm hedges, wasteland, rough pasture, modified lowland forest. More vigorous in deeper valley soils (Arulambalam 1998). Today, Japanese honeysuckle is widespread in the northern South Island but uncommon in the south part of the island. On the North Island, it is currently listed as widespread in almost all conservancies (Owen 1997). Typical areas invaded in New Japanese honeysuckle ( Lonicera japonica) 4 Zealand include the hilly landscapes where there are untrimmed roadsides and unattended land and associated areas of shrubbery, scrub, and forest remnants (Williams and Timmins 1997). In these areas, Japanese honeysuckle is described as "patchy" and displays a distributional pattern similar to other weedy plants whose primary dispersal agent is humans, with secondary spread occurring by birds and other animals. After a gradual secondary spread, plants become established then spread vegetatively, and so on (Williams and Timmins 1997). Japanese honeysuckle still has much room to grow in New Zealand. Making it illegal for humans to disperse the plant may retard its invasion to areas still free of the weed. In North America, Japanese honeysuckle was introduced in 1806. By the 1860's, it was widely cultivated. It was first noted from the wild in 1882. The first documentation of its spread was in 1904, almost one hundred years after its arrival (Andrews 1919; Leatherman 1955). Today, Japanese honeysuckle is widespread throughout the eastern United States. It is listed as a noxious weed in Illinois and as a pest plant in several other eastern states. Climate where invading: Cool, mesic to wet areas. Zones 4-10 (Turner 1997). Japanese honeysuckle readily invades open natural communities, often by seed spread by birds. An aggressive colonizer of successional fields, this vine also will invade mature forest and open woodlands. Forests with either natural or unnatural openings are often invaded by Japanese honeysuckle when birds drop seeds into these light gap areas. Deep shading reduces the amount of invasion (Nyboer 1990). On Kauai, Japanese honeysuckle is escaping and spreading in mesic to wet areas in Koke'e State Park, and near Volcano and slopes of Hualalai, Hawai'i (Wagner et al. 1990). On Maui, Japanese honeysuckle can be seen planted widely in a variety of conditions. It has been successfully planted from sea level up to 4,200 ft (1,280 m), in dry and wet areas, in shade and full sun, and in a variety of soils. Although, few naturalized or wild populations can be found to date on Maui. Most naturalized populations have spread vegetatively from plantings. Two naturalized populations are located below Haleakala National Park, on Crater Road, Kula at 4,000 ft (1,219 m) and 4,200 ft (1,280 m) elevation. The average annual rainfall in this area is about 40 in (Juvik and Juvik 1998). One naturalized population is in full sun, possibly spread vegetatively from a nearby planting. The soil is loamy and well drained. The other is in the full shade growing in a grove of Eucalyptus, possibly spread by bird dispersal, as there are no signs of an intentional planting, though this may be an old house site or abandoned water tank, suggesting the plants could have been planted and are persistent. The soil is similar to that of the first population mentioned. Another naturalized population is at the Waikapu golf course, located at about 800 ft (244 m) elevation, spreading vegetatively into the unmaintained surrounding scrub. Average annual rainfall at this site is about 30 in (Juvik and Juvik 1998). In North America, Japanese honeysuckle grows up to 5,905 ft (1,800 m) in both open and shaded areas (Williams and Timmins 1997). Climatic limitations to growth in North America include frosts in the northern region which causes death of shoots, low Japanese honeysuckle ( Lonicera japonica) 5 precipitation in the western region, and lack of cold temperatures to break seed dormancy in the south (Leatherman 1955). In New Zealand, plants grow from sea level up to 2,438 ft (743 m) (Williams and Timmins 1997). It is one of the few species of plants that are tolerant of pollution from heavy metals and SO2 (Caiazza and Quinn 1980). Value to humans: Useful as a ground cover or to quickly hide fences and posts. Some varieties have been used in Chinese medicine since the Tang dynasty in 659 AD (Turner 1997). Japanese honeysuckle has been widely cultivated as an ornamental plant, as a road bank stabilizer, and as food and shelter for wildlife (Handley 1945). Problems: Japanese honeysuckle is considered a major pest due to its ability to outcompete and shade out native vegetation (Cacek 1998). This aggressive vine seriously alters or destroys the understory and herbaceous layers of the communities it invades, including prairies, barrens, glades, flatwoods, savannas, floodplain and upland forests. Japanese honeysuckle also may alter understory bird populations in forest communities (Nyboer 1990). In New Zealand, Japanese honeysuckle infests forests and climbs over shrublands or low canopy trees, especially on forest margins. Honeysuckle may form a complete blanket over small trees and shrubs. It is becoming an increasingly troublesome species of shrubland, forest margins, and open roadsides (Arulambalam 1998). Japanese honeysuckle is toxic if large quantities are eaten. The vines have saponic and cyanogenic glycosoides. The fruits have carotenoids (Russell 1997). Effects from eating large quantities of the fruit include vomiting, diarrhea, pupil dilation, cold sweat, rapid heartbeat, respiratory failure, convulsion, and coma (Russell 1197). Japanese honeysuckle has been known to escape from gardens to become a serious weed elsewhere. In Hawaii, plants were previously thought to not bear fruit (Wagner et al. 1990). Wagner et al. (1990) suggest that if fertile strains develop, Japanese honeysuckle could become an extremely serious weed. Recently, fruit has been observed on both Oahu (Wagner et al. 1990) and on Maui. Viable fruit bearing plants allows dispersal by birds and other animals into areas far from parent plants, in a polka dot pattern, whereas non-fruit bearing plants can only spread vegetatively, in an amoebae like pattern. To date, most naturalized populations observed have spread vegetatively from plantings. Biology and ecology: Japanese honeysuckle climbs and drapes over native vegetation, shading it out. It is capable of completely covering herbaceous and understory plants, and climbing trees to the canopy. The semi-evergreen condition of this honeysuckle allows for growth both prior to and after dormancy of other deciduous plants. The prolific growth covers and smothers vegetation present including understory shrubs and trees in forested communities. Vegetative runners are most prolific in the open sun and will resprout where touching the soil, forming mats of new plants. This honeysuckle will display little growth under moderate shade. In deep shade, runners develop but often die back. Flowering and seed development are heaviest in open-sun areas. Seedling establishment and growth is slow in the first 2 years of development of a new honeysuckle colony (Nyboer 1990). Japanese honeysuckle ( Lonicera japonica) 6 Associated animals: Insects known to feed on Japanese honeysuckle in North America include indigenous hawk moths (Sphingidae), wax moths (Gelechiidae) and bumble bees have been observed extracting pollen from flowers (Williams and Timmins 1997). A wide range of birds are known to feed on fruit in North America from turkeys to small passerines (Williams and Timmins 1997). Because of the high water content in the fruit, the seeds pass quickly through birds (Handley 1945). Fruits are also eaten by mammals such as deer and possums. Plants respond rapidly to herbivory pressures. However, Japanese honeysuckle spread rapidly after forest reserves in New Zealand were fenced and animals excluded (Williams and Timmins 1997). Seed viability: According to Wagner et al. (1990), Japanese honeysuckle does not usually produce fruit in Hawaii and if a fertile strain were to develop, it could become an extremely serious weed. Recently, fruit bearing plants have been observed on Oahu (Wagner et al. 1990). Japanese honeysuckle does indeed produce fruit on Maui as well. Several locations of Japanese honeysuckle populations were monitored throughout the year for phenology data and fruit production. The site descriptions are as follows. Costco, located in Kahului, at sea level has several plantings surrounding the property using Japanese honeysuckle as a ground cover, where it is spreading vegetatively into unmaintained adjacent lots. Waikapu golf course, located in Waikapu, at about 800 ft (244 m) on West Maui, also has several plantings escaping vegetatively into adjacent unmaintained lands. Eddy Tam Park, located in Makawao, at 1,600 ft (488 m) elevation, has Japanese honeysuckle planted as a ground cover on steep banks. Another site is a residential planting located on Kekaulike Ave., Kula, at 3,700 ft (1,128 m) elevation where Japanese honeysuckle is escaping into the adjacent gulch. The last two sites are located on Crater Rd., Kula, one at 4,000 ft (1,219 m) and another just above at 4,200 ft (1,280 m). At these two sites, the Japanese honeysuckle seemed slightly different in form and fruit from other populations. We distinguish the two by the latter having a twining habit, lighter green leaves, and glaucous, greenish-grey fruit. The different form may prove to be a different species or variety. There are 180 different species of Lonicera and several different cultivars of Lonicera japonica. Or, the differences may be attributed to climatic factors. Further investigation is needed to be sure. Fruit has been found at all sites checked to date. Fruit was first observed on December 15, 1998 on Kekaulike Avenue, Kula. Both immature and mature fruit was observed. Immature fruit was bright green and glossy. Mature fruit was slightly larger, shiny, and black. Fruit was observed at the Costco site on December 16, 1998. Most fruit found were immature, small, shiny, and green. Fruit was abundant and found in clusters of four. Fruit was observed at the 4,000 ft elevation site on December 19, 1998. Most fruit at this site was immature. Both immature and ripe fruit was slightly different in color than fruit observed at other sites, appearing a bluish-grey, green color, with a waxy white film (glaucous). Fruit was found at the 4,200 ft Eucalyptus grove site on January 5, 1999. Fruit appeared similar to that found at the 4,000 ft site. Fruit was found at the Eddy Tam site also on January 5, 1999. Mature fruit was black, and glossy, similar to typical fruit found. Once fruit was observed on Maui, the next question to answer was whether or not the seeds in the fruit were viable. Fruit was collected from the 4,000 ft site on January 5, Japanese honeysuckle ( Lonicera japonica) 7 1999 and from the Kekaulike Ave. site on December 15, 1998 for further comparisons and germination tests. Site descriptions: Kekaulike Ave.: Japanese honeysuckle appeared planted on a bank bordering Kekaulike Ave. Plants were naturalizing into Keahuaiwi gulch, by Pulehu bridge, along with other weedy horticulture plants such as Mexican daisy (Erigeron karvinskianus), cape ivy (Senecio mikanioides), Vinca (Vinca spp.), Cotoneaster (Cotoneaster spp.), and butterfly bush (Buddleia madagascariensis). This population is representative of common Japanese honeysuckle. 4,000 ft: Japanese honeysuckle is located on the side of the road, climbing up a telephone pole wire. It may be spreading from a planting, however, this in inconclusive. It is mixed with other weedy plants such as hill raspberry (Rubus niveus), waddle (Acacia mearnsii), olive (Olea europaea cuspidata), and cotoneaster (Cotoneaster spp.). This form appears different than common Japanese honeysuckle. Fruit descriptions: Kekaulike Ave.: Ripe fruits were shiny black, measuring from 5 mm to 10 mm in diameter. 46 fruits were collected. One fruit had a sizeable hole with seed aril and seeds still intact. Pulp from fruit was greenish and stained paper on which it was a greenish color. Another fruit found did not have any seeds or pulp left as if eaten out by some thing. It is not known what may have eaten the seeds and pulp and left the skin of the fruit. 4,000 ft: The fruit has a thing skin, like that of a grape. It starts out green and turns black to purple as the fruit ripens. There is a sort of white haze on the surface or skin. The pulp is green and the seeds are green to brown. The fruit looks smaller than other Japanese honeysuckle fruit we have observed at Kula, Kahului, and Makawao. We had always thought this Japanese honeysuckle looked different, with its viney habit, leaves that look a little lighter in color, more fruit per axil, and fruit are smaller with a white sheen. There are also less seeds per fruit. The following table displays these differences in fruit characteristics. Site Kekaulike Ave. 4,000 ft Ave. diameter of fruit (mm) 5.78 4.79 Ave. # seeds per fruit 5.3 1.56 Ave. diameter of seed (mm) 4.38 3.075 Germination: After seeds were removed from the fruit, 50 random seeds from each site were placed in petri dishes, 25 seeds per petri dish, on moist paper towels. Seeds were kept moist by periodically adding bottled water to the petri dishes. Seeds were monitored for germination. Germination was defined as sending out a root radicle. Germination percentages may increase further over time, however, for the purposes of this report, the experiment ran for 10 weeks. Japanese honeysuckle ( Lonicera japonica) 8 Out of the 50 seeds from the Kekaulike site, one germinated, or 2 %, after 10 weeks. Out of the 50 seeds from the 4,000 ft site, 27 germinated, about 54 %, after 10 weeks. This raises a few questions. Is the 4,000 ft site population a different strain that is more viable and more likely to invade in Hawaii? Or, are there climatic factors such as high elevation and colder temperatures, which contribute to greater seed viability at this site? Further investigation is needed to make these conclusions. Japanese honeysuckle was first found naturalizing in Hawaii in 1951 (Wagner et al. 1990). It is a serious weed in eastern North America and New Zealand where it is now illegal to sell, propagate, distribute, or commercially display it. On Maui, Japanese honeysuckle is currently being planted widely in many areas where it has spread vegetatively into adjacent lands. Viable fruit producing plants will allow long distance dispersal to lands currently free of Japanese honeysuckle. Public awareness and further documentation of its invasive characteristics are needed in order to decrease further plantings and spread of Japanese honeysuckle. Physical control: Cutting: Vines may be chopped just above ground level. Cutting is repeated every two weeks to deplete nutrient reserves in the roots and prevent resumption of photosynthesis. Cutting does not affect roots, which will continue to grow until their energy and nutrient supplies are depleted (Cacek 1998). Mowing limits the length of Japanese honeysuckle vines, but will increase the number of stems produced (Nyboer 1990). Flaming: By placing a kerosene torch over leaves on the same schedule as cutting, foliage is wilted and nutrient supplies in the roots are depleted. As with cutting, flaming will not affect roots. Burning: Although few quantitative studies occur in the literature, Barden and Matthews (1980) recommend controlled burning. Two annual burns in an experimental plot reduced honeysuckle crown volume by 80%. Ground cover was reduced by 35%. Fires killed most aboveground vines, but ground cover was maintained by re- sprouting roots. Burning may be combined with previous flaming, which wilts and dries leaves, providing fuel for the burn (Cacek 1998). Grubbing: Consists of mechanical removal and destruction of the entire plant, including the root. If all root tissue is removed, no regrowth can occur, and repetition is not necessary. Grubbing is labor intensive and may be locally destructive. Grubbing is most effective from fruiting to winter and early spring when plant reserves are lowest. Grazing: Controlled grazing by goats may serve to reduce honeysuckle crown and ground cover densities, but as with controlled burning, re-sprouting roots will regenerate unless nutrient reserves are depleted by continuous grazing pressure. Regardless of the control method used, care must be taken to prevent re- invasion from nearby areas, or by seeds transported by birds or other wildlife. Planting the area with fast-growing native vegetation or grasses may prevent recolonization (Cacek 1998). Japanese honeysuckle ( Lonicera japonica) 9 Chemical control: Mclemore (1981) reports that an acceptable level of control (70%) was reached during a two-year experimental program which used 2 lb/acre of glyphosate in the first year and 6 lb/acre in the second year. Japanese honeysuckle is an evergreen, so it can be treated in the dormant season with less damage to non-target species (Cacek 1998). In New Zealand, they cut base of stem and paint with stump paint herbicide mix. Alternatively spray with Escort + Pulse or Tordon Brushkiller herbicide. Cut large vines at head height and spray below this point. Roundup has been found to be ineffective (Arulambalam 1998). Spray infestations on open ground with Escort. To save trees which are growing under or through honeysuckle, cut and paint stems or use Tordon Granules (Arulambalam 1998). In temperate areas, because Japanese honeysuckle is semi-evergreen, it will continue to photosynthesize after surrounding deciduous vegetation is dormant. This condition allows managers to detect the amount of infestation, and allows for treatment of the infestation with herbicides without damage to the dormant vegetation (Nyboer 1990). Glyphosate herbicide (trade name Roundup) is the recommended treatment for this honeysuckle. A 1.5-2% solution (2-2.6 oz Roundup/gallon water) applied as a spray to the foliage will effectively eradicate Japanese honeysuckle. Although glyphosate is effective when used during the growing season, use at this time is not recommended in natural areas because of the potential harm to nontarget plants (Arulambalam 1998). Crossbow, a formulation of triclopyr and 2,4-D, is also a very effective herbicide that controls Japanese honeysuckle. Crossbow should be mixed and applied as a foliar spray. Herbicides that have given poor control results or that are more persistent in the environment than other types are picloram, annitrole, aminotriazole, atrazine, dicamba & 2,4-D, 2,4_D, DPX 5648, fenac, fenuron, simazine & triclopyr. Biological control: No natural enemies are reported for this species (Cacek 1998). Management recommendations: Japanese honeysuckle has a extensive history as an invasive species, yet is widely planted on Maui in schools, parks and housing complexes and can be bought in most garden shops. Recently, viable fruit have been found, and it has been seen spreading into wild lands. Perhaps the best strategy for Japanese honeysuckle is to stop selling it and control infestations adjacent to wild lands. Raising public awareness may help to prevent future plantings, and putting it on the State Noxious Weed List would provide legal authority for those looking to control the spread of Japanese honeysuckle. Key contacts: Maui: USGS/BRD Haleakala Field Station PO Box 369 Makawao, HI 96768 (808) 572-4472 Kaua’i: Japanese honeysuckle ( Lonicera japonica) 10 Guy Nagai Hawai’i State Department of Agriculture 4398A Pualoke St. Lihue, HI 96766-1673 (808) 274-3069 REFERENCES & BIBLIOGRAPHY: Andrews, E.F. 1919. The Japanese honeysuckle in the eastern United States. Torreya 19: 37-43. Aukland Regional Council. 1997. Pest Facts #43, Japanese honeysuckle, Lonicera japonica. Auckland. Arulambalam, R. 1998. Weeds of New Zealand. http://www.boprc.govt.nz/www/green/WEED63.HTM. Barden, L. S., and J. F. Matthews. 1980. Change in abundance of honeysuckle (Lonicera japonica) and other ground flora after prescribed burning of a piedmont pine forest. Castanea 45:257-260. Cacek, T. 1998. The National Park Service Integrated Pest Management Manual. http://www.colostate.edu/Depts/IPMnatparks/exweeds2.html. Caiazza, N.A., and J.A. Quinn. 1980. Leaf morphology in Arenaria patula and Lonicera japonica along a pollution gradient. Bulletin of the Torrey Botanical Club 107: 9-18. Carr, G.W., J.V. Yugovic, and K.E. Robinson. 1992. Environmental weed invasions in Victoria. Department of Conservation and Environment and Ecological Horticultural Pty. Ltd., Melbourne. 78 p. Craw, J. 1994. Keeping our gardens forest friendly. Forest and Bird 271: 8-12. Cronk, Q.C., and J. Fuller. 1995. Plant Invaders. Chapman Hall, London. 241 p. Esler, A.E. 1988. Naturalised plants in urban Auckland: a series of articles from the New Zealand Journal of botany. DSIR Publishing, Wellington. 618 p. Handley, C.O. 1945. Japanese honeysuckle in wildlife management. Journal of Wildlife Management 9: 261-264. Juvik S.P and J.O. Juvik. 1998. Atlas of Hawai'i. Third edition. Department of Geography, University of Hawai'i press. Leatherman, A.D. 1955. Ecological life-history of Lonicera japonica Thunb. Unpublished PhD thesis, University of Tennessee (Library of Congress Card No. Mic. Japanese honeysuckle ( Lonicera japonica) 11 55-772). 97 p. University Microfilms, Ann Arbor, Michigan (Dissertation Abstracts 15 (11): 1987, Publication No. 15,076). McLemore, B.F. 1982. Comparison of herbicides for controlling hardwoods in pine stands. Proceedings of the 35th annual meeting of the Southern Weed Science Society 1982. 195-199. Nyboer, R. 1990. Vegetation Management Guideline Japanese Honeysuckle. http://www.inhs.uiuc.edu/edu/VGM/jhnysckl.html. Owen, S.J. 1997. Ecological weeds on conservation land in New Zealand: A database. Department of Conservation, Wellington. Russell, A. B. 1997. Poisonous Plants of North Carolina. http://russell4.hort.ncsu.edu/poison/Lonicja.htm. Turner, R. J. Jr., E. Wasson. 1997. Botanica. Random House Australia. NSW Australia. 1007 pp. University of Hawaii, Botany Department Web Page. Alien plants of Hawaii. Lonicera japonica. http://www.botany.hawaii.edu/faculty/cw_smith/lon_jap.htm. Wagner, W.H., D.R. Herbst, S.H. Sohmer. 1990. Manual of the Flowering Plants of Hawai’i. University of Hawai’i press, Honolulu, Hawai’i. Williams, P.A., and S.M. Timmins. 1997. Biology and ecology of Japanese honeysuckle (Lonicera japonica) and its impacts in New Zealand. Landcare Research Contract Report: LC 9798/013. D Japanese honeysuckle ( Lonicera japonica) 12 Japanese honeysuckle fruit and flower Japanese honeysuckle rooting at the nodes - Kula, Maui Japanese honeysuckle ( Lonicera japonica) 13 Typical use of Japanese honeysuckle in landscaping - Eddie Tam Park, Makawao, Maui A stand of Japanese honeysuckle escaping into Eucalyptus grove - Kula, Maui Japanese honeysuckle ( Lonicera japonica) 14 Japanese honeysuckle fruit used in germination trials. Counting Japanese honeysuckle seeds and preparing them for germination trials Japanese honeysuckle ( Lonicera japonica) 15 Known global distribution of Japanese honeysuckle Known distribution of Japanese honeysuckle in the state of Hawaii, USA Japanese honeysuckle ( Lonicera japonica) 16 Known distribution of Japanese honeysuckle on the island of Maui, state of Hawai'i Japanese honeysuckle ( Lonicera japonica) 17
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